Volume 131, number 2 FEBS LETTERS August 1981
CALCIUM AND CALMODULIN-DEPENDENT PROTEIN PHOSPHORYLATION IN RABBIT
ILEUM
Leslie TAYLOR, Victoria J. GUERINA, Mark DONOWITZ, Michael COHEN and Geoffrey W. G. SHARP*
Departments of Physiology and Medicine, Tufts University School of Medicine and The New England Medical Center Hospital
Boston, MA 02111, USA
Received 13 July 1981
1. Introduction
Evidence is accumulating to suggest that calcium is
a physiological regulator of intestinal electrolyte
transport [ 1-3]. Conditions which increase intracel-
lular calcium such as the use of the calcium ionophore
A23187 [ 1,2] or exposure to neurohumoral sub-
stances such as serotonin [4] or carbachol [ 1] cause
stimulation of intestinal chloride secretion and/or
inhibition ofNa ÷and C1- absorption. To the contrary,
conditions which decrease intracellular calcium, such
as exposure to the calcium channel blocker verapamil,
stimulate Na ÷ and C1- absorption [3]. Calmodulin
may be involved in these effects since the anti-psy-
chotic drug trifluoperazine (an inhibitor of the cal-
cium-calmodulin complex) inhibited intestinal secre-
tion caused by the calcium ionophore A23187 [5,6].
The intracellular mechanisms by which calcium
and calmodulin affect intestinal ion transport are not
known. However, as calcium and calmodulin affect
phosphorylation and function of specific proteins in
several other systems [7,8], phosphorylation is a
potential control mechanism for ileal electrolyte
transport. These studies are the first demonstration
that calcium and calmodulin can cause phosphoryla-
tion of intestinal peptides.
2. Materials and methods
Fed white male New Zealand rabbits (2-2.5 kg
* Present address: Department of Pharmacology,New York
State College of Veterinary Medicine, Cornell University,
Ithaca, NY 14853, USA
Address correspondence and reprint requests to: Dr Mark
Donowitz, GastroenterologyUnit, Tufts-New England Medi-
cal Center, 171 Harrison Avenue, Boston, MA 02111, USA
body wt) were anesthetized with sodium pentobarbital
and the distal ileum removed. After washing with
cold Ringer's-HCO3 solution, mucosal scrapings were
obtained on ice using glass slides. This technique has
been demonstrated to obtain primarily villous tip
cells and some crypt cells [9]. 150 mg tissue was
homogenized in 7.5 ml 5 mM MgC12, 10% sucrose,
50 mM Tris-HC1 (pH 7.5) using a ground-glass hand
homogenizer. The homogenate, containing ~200/2g
protein, was preincubated in a 100/21 reaction mix-
ture containing 1 mM EGTA, 5 mM MgC12, 10%
sucrose, 50 mM Tris-HC1 (pH 7.5) and, where indi-
cated, 1.1 mM CaC12, and different concentrations of
calmodulin and trifluoperazine, for 1 min at 37°C.
The phosphorylation reaction was started by the
addition of [32p] ATP at a final concentration of 5/2M.
The reaction was stopped after 20 s by the addition
of 50/21 of a solution containing 0.1 mM EDTA, 5%
SDS, 200 mM dithiothreitol and 50 mg/ml pyronin
Y followed immediately by immersion in a boiling
water bath for 2 min. The mixture was then placed
in a 37°C water bath for 20 rain. Samples, contain-
ing --40/2g protein, were subjected to electrophoresis
on a polyacrylamide continuous gradient slab gel
(5-15% acrylamide) according to [10]. The gels
were subjected to autoradiography using XAR-5
Kodak film (Eastman Kodak, Rochester NY) with
exposure times ranging from 4-48 h. The autoradio-
graphs were subsequently analyzed by use of a Zeneih
scanning densitometer (Biomed Products, Chicago IL).
Amount of phosphorylation was assumed propor-
tional to peak height of the densitometry scan [ 11 ].
Between 6 and 8 experiments were performed for
each experimental condition.
[TP2P] Adenosine triphosphate (spec. act. 3-10 Ci/
mmol) was obtained from New England Nuclear
Published by Elsevier/North-Holland Biomedical Press
322 00145793/81/0000-0000/$02.50 © 1981 Federation of European BiochemicalSocieties